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The Human Career: Human Biological and Cultural Origins, Third Edition
The Human Career: Human Biological and Cultural Origins, Third Edition
The Human Career: Human Biological and Cultural Origins, Third Edition
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The Human Career: Human Biological and Cultural Origins, Third Edition

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Since its publication in 1989, The Human Career has proved to be an indispensable tool in teaching human origins. This substantially revised third edition retains Richard G. Klein’s innovative approach while showing how cumulative discoveries and analyses over the past ten years have significantly refined our knowledge of human evolution.

Klein chronicles the evolution of people from the earliest primates through the emergence of fully modern humans within the past 200,000 years. His comprehensive treatment stresses recent advances in knowledge, including, for example, ever more abundant evidence that fully modern humans originated in Africa and spread from there, replacing the Neanderthals in Europe and equally archaic people in Asia. With its coverage of both the fossil record and the archaeological record over the 2.5 million years for which both are available, The Human Career demonstrates that human morphology and behavior evolved together. Throughout the book, Klein presents evidence for alternative points of view, but does not hesitate to make his own position clear.

In addition to outlining the broad pattern of human evolution, The Human Career details the kinds of data that support it. For the third edition, Klein has added numerous tables and a fresh citation system designed to enhance readability, especially for students. He has also included more than fifty new illustrations to help lay readers grasp the fossils, artifacts, and other discoveries on which specialists rely. With abundant references and hundreds of images, charts, and diagrams, this new edition is unparalleled in its usefulness for teaching human evolution.

LanguageEnglish
Release dateApr 22, 2009
ISBN9780226027524
The Human Career: Human Biological and Cultural Origins, Third Edition

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  • Rating: 4 out of 5 stars
    4/5
    This 1989 edition has been updated in 2000. I have not yet acquired the newer version, but even this is worthwhile for getting your feet wet on the subject of human evolution. Klein is a careful scholar, weighing evidence clearly and letting you know on what grounds he has come to his conclusions. He lays out evidence very well, and that alone makes this recommended reading, because whatever else you read on the subject, you have to understand the kinds of evidence that scientists use and how they come to conclusions from that evidence. Since this topic is a hot one, new discoveries are occurring all the time and older evidence is being reevaluated. Even so, I found Klein's chronicalling of the development of hominids into modern humans, very enlightening and not contradictory of newer evidence that I have read.

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The Human Career - Richard G. Klein

Richard G. Klein is professor of anthropology and biology at Stanford University. His books include Ice Age Hunters of the Ukraine and (with Kathryn Cruz-Uribe) The Analysis of Animal Bones from Archeological Sites, both published by the University of Chicago Press.

The University of Chicago Press, Chicago 60637

The University of Chicago Press, Ltd., London

© 1989, 1999, 2009 by The University of Chicago

All rights reserved. Published 2009

Printed in the United States of America

18 17 16 15 14 13 12 11 10 09       1 2 3 4 5

ISBN-13: 978-0-226-02752-4 (e-book)

ISBN-13: 978-0-226-43965-5 (cloth)

ISBN-10: 0-226-43965-8 (cloth)

Library of Congress Cataloging-in-Publication Data

Klein, Richard G.

The human career : human biological and cultural origins / Richard G. Klein.—3rd ed.

p. cm.

Includes bibliographical references and index.

ISBN-13: 978-0-226-43965-5 (cloth: alk. paper)

ISBN-10: 0-226-43965-8 (cloth: alk. paper)

ISBN 978-0-226-02752-4 (e-book)

1. Human beings—Origin.   2. Fossil hominids.   3. Human evolution.   I. Title.

GN281.K55 2009

599.93' 8—dc22

2008029270

The paper used in this publication meets the minimum requirements of the American National Standard for Information Sciences—Permanence of Paper for Printed Library Materials, ANSI Z39.48-1992.

THE HUMAN CAREER

Human Biological and Cultural Origins

THIRD EDITION

Richard G. Klein

THE UNIVERSITY OF CHICAGO PRESS

Chicago and London

CONTENTS

List of Illustrations

List of Tables

Preface to the Third Edition

Preface to the Second Edition

Preface to the First Edition

1. Evolution, Classification, and Nomenclature

2. The Geologic Time Frame

3. The Primate Background

4. The Australopiths and Homo habilis

5. Evolution of the Genus Homo

6. The Neanderthals and Their Contemporaries

7. Anatomically Modern Humans

8. Synopsis: Anatomy, Behavior, and Modern Human Origins

References

Reference Index

Site Index

Subject Index

ILLUSTRATIONS

1.1. The process of speciation as visualized by gradualists and by advocates of punctuated equilibrium

1.2. A currently popular phylogeny of the hominins

1.3. A popular hierarchical classification of the living Primates down to the family level

1.4. A cladogram illustrating the evolutionary relationships among humans, chimpanzees, gorillas, orangutans, gibbons, rhesus monkeys, and spider monkeys

2.1. The geologic timescale and the proposed ages of some important evolutionary events

2.2. Stages in the evolution of Mimomys, Arvicola, and Microtus

2.3. Middle and late Pleistocene vole stratigraphy of central and southeastern Europe

2.4. Shed mature antlers of red deer from Mosbach, Germany, and Ilford, England

2.5. The average number of plates on third molars in successive species of Mammuthus

2.6. Time ranges of the Elephantinae in Africa

2.7. Time ranges of fossil suids in Africa after 10 Ma

2.8. Time ranges of the later Miocene to Holocene equids in Africa

2.9. Time ranges covered by various numerical dating methods

2.10. A proposed curve for calibrating radiocarbon ages to calendar (solar) years

2.11. Global paleomagnetic stratigraphy for the past 5 million years

2.12. Global temperature change over the past 70 million years

2.13. The δ oxygen-18 record for the past 900,000 years

2.14. The maximum extent of Quaternary glaciation and associated sea-level change

3.1. Reconstructed skull of Australopithecus africanus

3.2. Right upper and lower dentitions of various primates

3.3. Reconstructed skeleton of Australopithecus afarensis

3.4. Diagrammatic section through the right ear region of a therian mammal

3.5. Geographic distribution of the living nonhuman primates

3.6. Nostril orientation in New World and Old World monkeys

3.7. Cephalic views of the rib cage and right shoulder girdle of an adult macaque monkey and of an adult human

3.8. Lower molars of Propliopithecus and Colobus

3.9. Occlusal relation between the upper canine, lower canine, and lower third premolar in a nonhuman catarrhine primate and a human being

3.10. Quadrupedal postures in an Old World monkey, a human, a gorilla, and a chimpanzee

3.11. Skulls of male and female adult mandrills

3.12. Skulls of a Malagasy lemur and a New World monkey

3.13. Biomolecular estimates for the divergence times between great apes, Old World monkeys, New World monkeys, and strepsirrhines

3.14. Changing positions of the continents from the late Triassic to the middle Eocene

3.15. Two interpretations of early primate phylogeny

3.16. Skulls, foot bones, and reconstructions of Paleocene and Eocene primates

3.17. Eocene and Oligocene localities with possible or probable higher primates fossils

3.18. Mandibles of Apidium phiomense, Propliopithecus chirobates, and P. zeuxis

3.19. Facial and lateral views of the skull of Propliopithecus zeuxis

3.20. Anterior and distal views of the distal left humerus in various monkeys and in Propliopithecus zeuxis

3.21. Temporal distribution of fossil hominoid genera

3.22. Relative positions of Africa and Eurasia in the early and middle Miocene

3.23. Approximate locations of the main African Miocene fossil hominoid sites

3.24. Reconstructed skull and skeleton of Proconsul heseloni

3.25. Fossil hominoid localities dating between about 16–15 and 8–7 Ma in relation to the historic distribution of the chimpanzees and the gorilla

3.26. Reconstructed skeleton of Pliopithecus vindobonensis

3.27. Skulls of Sivapithecus indicus, Pongo pygmaeus, and Pan troglodytes

3.28. A provisional phylogeny of the Primates

4.1. Approximate locations of African Plio-Pleistocene fossil sites

4.2. Facial skeleton and endocast of Australopithecus africanus from Taung

4.3. The position of the skull relative to the spinal column in a gorilla and a modern human

4.4. Ages of some important South African hominin fossil sites, inferred from mammalian species

4.5. Dating of the earliest artifact industries in Africa and Eurasia

4.6. Time spans of the most commonly recognized hominin species between 4.4 and 1.0 million years ago

4.7. Three stages in the evolution of the Swartkrans australopith cave

4.8. The location of Olduvai Gorge on the Serengeti Plain

4.9. Schematic stratigraphy of Olduvai Gorge

4.10. Schematic stratigraphic column for the Laetoli area

4.11. Distribution of fossiliferous geological formations in the Lake Turkana Basin

4.12. Schematic stratigraphy of the Koobi Fora Formation and correlation to the Shungura Formation

4.13. The approximate numbers of hominin fossils by member within the principal formations of the Omo Group

4.14. Correlation of the Plio-Pleistocene successions in the Lake Turkana Basin

4.15. Schematic stratigraphy of the Hadar Formation

4.16. The skull of Sahelanthropus tchadensis before reconstruction

4.17. Posterior view of the Orrorin tugensis proximal femur

4.18. Insertion of the masseter and temporalis muscles on a modern human skull

4.19. Mandibular dentitions of a female chimpanzee, Ardipithecus ramidus, and Australopithecus afarensis

4.20. Upper canines and lower premolars of Dryopithecus, male and female chimpanzees, Ardipithecus ramidus, and Australopithecus afarensis

4.21. Lower deciduous anterior premolars of Dryopithecus, pygmy chimpanzee, common chimpanzee, Ardipithecus ramidus, Australopithecus afarensis, Au. africanus, Paranthropus robustus, P. boisei, and living humans

4.22. Schematic views of the skull of Australopithecus afarensis

4.23. Basal views of the skull of Pan troglodytes, Australopithecus afarensis, A. africanus, Paranthropus robustus, and modern Homo sapiens

4.24. Occlusal views of mandibles of Australopithecus anamensis and A. afarensis

4.25. Lingual views of mandibles of Australopithecus anamensis and A. afarensis

4.26. Proximal tibiae of a chimpanzee, a living human, and Australopithecus anamensis

4.27. Lateral and anterior views of cranium AL 444-2, Australopithecus afarensis from Hadar

4.28. Facial and occipital views of Pan troglodytes, Australopithecus afarensis, A. africanus, Paranthropus robustus, P. boisei, and Homo habilis

4.29. Palates of a chimpanzee, various australopiths, and a modern human

4.30. Upper canine morphology of various australopiths, chimpanzees, and modern humans

4.31. Cross sections of the mandibular body below the fourth premolar in various australopiths and in Homo habilis

4.32. Lower third premolar morphology in chimpanzees, in Australopithecus afarensis, and in modern humans

4.33. The superior margin of the mandibular ramus in Australopithecus afarensis, Gorilla gorilla, Paranthropus robustus, Pan troglodytes, and modern Homo sapiens

4.34. Occipital views of cranial venous sinus systems typical for modern humans and most other hominoids and for Australopithecus afarensis, Paranthropus boisei, and P. robustus

4.35. Thumb metacarpals of a modern human, Paranthropus robustus, Australopithecus afarensis, and a chimpanzee

4.36. Lower limb and knee joint of a modern human, Australopithecus afarensis, and a chimpanzee

4.37. Skulls and pelvises of a chimpanzee, Australopithecus afarensis, and a living human

4.38. Facial views of KNM-WT 40000 (Kenyanthropus platyops) and KNM-ER 1470 (Homo rudolfensis)

4.39. Basicranial flexion in gracile australopiths, robust australopiths, Homo habilis, and modern H. sapiens

4.40. Superior and lateral views of the skull of Australopithecus garhi

4.41. Maxillae of Australopithecus garhi and Paranthropus boisei

4.42. Relative limb bone proportions in Pan troglodytes, Australopithecus afarensis, possible Australopithecus garhi, Homo ergaster, and Homo sapiens

4.43. Skulls of Paranthropus aethiopicus and P. boisei

4.44. A mandible of Paranthropus robustus articulated with a reconstructed skull of P. boisei

4.45. A reconstructed male skull of Paranthropus robustus

4.46. Reconstructed skulls of Homo habilis from Koobi Fora

4.47. Palatal view of maxilla AL 666-1 (Homo cf. habilis) from Hadar

4.48. Cladograms showing the evolutionary relationships of the australopiths and Homo suggested by endocranial capacity and by cheek tooth area

4.49. A working phylogeny of the hominins

4.50. The basic types of stone artifacts found at Oldowan and Acheulean sites in Africa

4.51. A range of typical Oldowan stone tools and their conventional typological designations

4.52. Oldowan artifacts from sites EG10 and EG12, East Gona, Ethiopia

4.53. The relation between handedness and flake form

4.54. A partial floor plan of site DK 1 at Olduvai Gorge

4.55. Fossilized hyena feces, a cut-marked bone, and various tooth-marked bones

4.56. The abundance of large mammals at the FLK Zinj site, Olduvai Gorge Bed I

5.1. The phylogeny of the genus Homo

5.2. The occupied world roughly 500,000 years ago

5.3. Principal fossiliferous localities in Java

5.4. The main Chinese sites with fossils of primitive Homo

5.5. Approximate ages of the main sites with fossils of Homo ergaster, H. erectus, early H. neanderthalensis, and early H. sapiens

5.6. Tentative correlation of global marine oxygen-isotope stages and key European human fossil sites

5.7. Franz Weidenreich’s restorations of Homo erectus skulls from Java and China

5.8. Skulls of Javan Homo erectus and a robust modern person

5.9. Front and side views of Ngandong skull XI and Sangiran Skull 17

5.10. Browridge form in Indonesian and Chinese Homo erectus

5.11. Mandibles of Indonesian Homo erectus and a robust modern person

5.12. Skull KNM-ER 3733 (Homo ergaster) from Koobi Fora, East Turkana

5.13. Skull of KNM-WT 15000 (Homo ergaster) from Nariokotome III, West Turkana

5.14. Reconstructed skeletons of Homo ergaster and Australopithecus afarensis

5.15. Fossils of Homo ergaster from Swartkrans Cave

5.16. Fossil skullcap from the Daka Member of the Bouri Formation, Middle Awash Valley

5.17. Skullcap of Olduvai Hominid 9 from Upper Bed II, Olduvai Gorge

5.18. Skullcap of archaic Homo from the Narmada Valley, India

5.19. Fossil skull from Swanscombe, England

5.20. Rear views of skulls of Homo erectus, early H. sapiens, modern H. sapiens, early H. neanderthalensis, and classic H. neanderthalensis

5.21. Skulls 4, 5, and 6 from the Sima de los Huesos, Atapuerca

5.22. Facial view of skull 5 from the Sima de los Huesos, Atapuerca

5.23. Mandibles AT 505 and AT 605 from the Sima de los Huesos, Atapuerca

5.24. Midsagittal sections through the occipital bones of Homo erectus, early H. neanderthalensis, and later H. neanderthalensis

5.25. Fossil skulls from Steinheim (Germany), Arago (France), and Petralona (Greece)

5.26. Fossil human mandible from Montmaurin, France

5.27. The principal fossil skull from Bodo, Middle Awash

5.28. Fossil skull from deposits of seasonal Lake Ndutu

5.29. Fossil skull from Kabwe, Zambia

5.30. Fossil skullcap from Elandsfontein, South Africa

5.31. Skull 1 from Jebel Irhoud, Morocco

5.32. Fossil skull from Singa, Sudan

5.33. Fossil skull from Florisbad, South Africa

5.34. Fossil skull from Dali, China

5.35. Fossil skull from Jinniushan, China

5.36. Fossil skull from Maba, China

5.37. Probable human dispersal routes from Africa superimposed on the locations of early archaeological or human fossil sites in Africa and Eurasia

5.38. A schematic section through the Dmanisi site, Georgia

5.39. Skull 2700 and mandible 2735 from Dmanisi, Georgia

5.40. Mandible KNM-ER 922 from East Turkana, Kenya, and mandible D211 from Dmanisi, Georgia

5.41. Skulls D2280 and D2282 from Dmanisi, Georgia

5.42. Skull 3444 and mandible 3900 from Dmanisi, Georgia

5.43. The approximate locations of European sites with early human fossils or artifacts

5.44. Reconstructed fossil skullcap from Ceprano, Italy

5.45. A juvenile maxilla from the Gran Dolina, Atapuerca

5.46. Bifaces from site TK, Bed II, Olduvai Gorge

5.47. The approximate locations of the main African Acheulean sites

5.48. Earlier and later Acheulean bifaces from the Bouri Formation, Middle Awash

5.49. Earlier and later Acheulean hand axes from Sterkfontein Cave and Kathu Pan

5.50. Earlier and later Acheulean cleavers from Sterkfontein Cave and Elandsfontein

5.51. Later Acheulean artifacts from Elandsfontein Cutting 10

5.52. Artifacts associated with Homo erectus at Zhoukoudian Locality 1

5.53. Locations of the main excavated early Paleolithic sites in western Asia

5.54. Acheulean bifaces from Torralba, Spain

5.55. Late Acheulean artifacts from southern England

5.56. Tayacian artifacts from La Caune de l’Arago, France

5.57. Bone and artifact concentrations at Bilzingsleben, Germany

5.58. Floor plan of an early Middle Paleolithic layer at Le Lazaret Cave, France

5.59. A flaked fragment of elephant bone from Bilzingsleben, Germany

5.60. Wooden throwing spears from Schöningen, Germany

5.61. An elephant tibia shaft fragment with scored lines from Bilzingsleben, Germany

5.62. A putative figurine from Berkehat Ram, Golan Heights, and the Venus of Lespugue

5.63. Proportionate representation of different large mammals at the Elandsfontein Acheulean site

5.64. Proportionate representation of bovid species at the Elandsfontein Acheulean site

5.65. Proportionate representation of carnivore species at the Elandsfontein Acheulean site

5.66. A schematic section through the Sima de los Huesos, Atapuerca

6.1. The European and west Asian realm of the Neanderthals

6.2. Approximate locations of European and north African Middle Paleolithic sites

6.3. Skullcap of Neanderthal 1 from Feldhofer Cave

6.4. Approximate locations of southwest Asian Middle and Upper Paleolithic sites

6.5. Derived features on Neanderthal skulls from La Ferrassie Cave (France) and Amud Cave (Israel)

6.6. Neanderthal skull from La Quina Cave (France) and the non-Neanderthal skull from Broken Hill (Zambia)

6.7. Adult male Neanderthal and modern human skeletons

6.8. Labyrinthine inner ear structure in pygmy chimpanzees, modern humans, and the La Ferrassie 1 Neanderthal

6.9. Shoveling in Neanderthal upper front teeth from Krapina, Croatia

6.10. The dentition of Shanidar Neanderthal skull 1, illustrating the retromolar space and labial wear

6.11. Taurodont upper cheek teeth from Krapina Cave, Croatia

6.12. Superior margin of the mandibular ramus in a recent human, near-modern humans, and Neanderthals

6.13. Configuration of the mandibular foramen in a modern human from Vogelherd, Germany, and a Neanderthal from La Chapelle-aux-Saints, France

6.14. Femurs of a Neanderthal and of a modern person

6.15. The axillary border of the scapula in Neanderthals and modern humans

6.16. Hand skeletons of a Neanderthal and an early-modern human

6.17. The shaft/neck angle on proximal femurs of a Neanderthal and a modern Euramerican male

6.18. Reconstructed physiques of a Neanderthal and of an early-modern European

6.19. Superior view of the reconstructed Neanderthal pelvis from Kebara Cave, Israel

6.20. Approximate locations of key north African Aterian and Mousterian/Middle Stone Age (MSA) sites

6.21. Approximate locations of east African Middle and Later Stone Age sites

6.22. Approximate locations of the main MSA sites in southern Africa

6.23. A fossil skull from Jebel Irhoud, Morocco, and a Neanderthal skull from Spy Cave, Belgium

6.24. Facial and lateral views of the near-modern human skull from Ngaloba, Tanzania

6.25. Near-modern human skull and mandible from Dar es Soltan 2, Morocco

6.26. The principal adult skull from Herto, Ethiopia

6.27. Skulls 1 and 2 from the Omo-Kibish Formation, Ethiopia

6.28. Occlusal and buccal views of mandibles 41815 and 16424 from Klasies River Main, South Africa

6.29. The reconstructed face and skull of a Neanderthal associated with Châtelperronian artifacts at Saint Césaire, France

6.30. Skull 9 from Qafzeh Cave, Israel

6.31. Partial reconstitution of a flat, discoidal core by refitting of flakes from Mousterian Site C, Maastricht-Belvédère, the Netherlands

6.32. Stages in the manufacture of a classic Levallois core

6.33. Two variants of the Levallois technique for producing elongated flakes or blades

6.34. The technique that Quina Mousterians often used to slice flakes from an elongated core

6.35. Basic Mousterian and MSA stone-tool types, as defined by François Bordes

6.36. Reduction of a simple convex sidescraper to other sidescraper types

6.37. Convergent sidescrapers from the Mousterian site of Biache-Saint-Vaast, France

6.38. Utilized Levalloiso-Mousterian artifacts from Kebara Cave, Israel

6.39. Schematic section through the deposits of Combe-Grenal Cave, France

6.40. Correlation between the archaeological sequences at Le Moustier and Combe-Grenal, France

6.41. Paleolithic cultural stratigraphy and chronology of the Levant

6.42. Aterian artifacts from Algeria

6.43. Howieson’s Poort artifacts from Nelson Bay Cave, South Africa

6.44. Barbed bone points from Katanda, Democratic Republic of the Congo

6.45. The putative Mousterian flute from Divje Babe Cave 1, Slovenia

6.46. Proposed tick shell beads and an incised red ocher fragment from Blombos Cave, South Africa

6.47. Floor plan and section through find level II at Ariendorf, Germany

6.48. Plan of Molodova I, level 4, Ukraine

6.49. Approximate locations of South African sites that illuminate Middle and Later Stone Age coastal ecology

6.50. The minimum numbers of eland, Cape buffalo, and bushpig in the Stone Age levels of Klasies River Cave 1, Nelson Bay Cave, Die Kelders Cave 1, and Byneskranskop Cave 1, South Africa

6.51. Mortality profiles of eland and buffalo in the MSA layers of Klasies River Cave 1

6.52. Distal humerus breadth in known-age fur seals and in fossil or subfossil fur seal samples from Stone Age and brown hyena sites on the coast of southern Africa

6.53. The maximum length of Cape turban shell opercula in Stone Age sites on the south coast of South Africa

6.54. The maximum length of granite limpet shells in Stone Age sites on the west coast of South Africa

6.55. Breadths of angulate tortoise distal humeri in Stone Age sites on the south coast of South Africa

6.56. Breadths of angulate tortoise distal humeri in Stone Age sites on the west coast of South Africa

6.57. Face of Shanidar 1, showing the crushed outer margin of the left orbit

6.58. Chronological arrangement of late Pleistocene cultural units and fossil human types in Africa and Eurasia

6.59. The distribution of the Aurignacian, Châtelperronian, Uluzzian, and Szeletian/Jerzmanowician Industries in Europe and western Asia roughly 37 ka

6.60. Stone and bone artifacts from the Châtelperronian layers of the Grotte du Renne, France

6.61. Plan and reconstruction of the Châtelperronian hut emplacement in level XI of the Grotte du Renne, France

6.62. Two proposed routes for basal Aurignacian dispersal across Europe

6.63. Skulls of individual 1 from Shanidar Cave, Iraq, and of individual 3 from Předmostí, Czech Republic

6.64. Bifacial leaf-shaped points from the upper deposits of Szeleta Cave, Hungary

6.65. Body proportions in the early-modern or near-modern Last Interglacial occupants of Israel and the Last Glacial Neanderthals of Europe

6.66. Artifacts that characterize different variants of the Early Upper Paleolithic in southwestern Asia

7.1. Reconstructed skulls of the Cro-Magnon 1 early-modern European and the La Chapelle-aux-Saints Neanderthal

7.2. Approximate locations of major western and central European Upper Paleolithic sites

7.3. Skull of an anatomically modern child from Kostenki XV, Russia

7.4. Skull of an anatomically modern adolescent from the Pataud Rockshelter, France

7.5. A phylogram based on complete mtDNA sequences from fifty-three geographically dispersed individuals

7.6. Hypothetical routes of modern human dispersal from Africa

7.7. Typical Upper Paleolithic tool types

7.8. Typical Aurignacian split-base bone points, pendants, and chipped stone artifacts

7.9. Gravettian stone artifacts

7.10. Solutrean artifacts from La Riera Cave, Spain

7.11. Magdalenian bone and stone artifacts from El Juyo Cave, Spain

7.12. Climate and cultural stratigraphy after 186,000 years ago

7.13. Abundance of mammal species in the Magdalenian layers of El Juyo Cave Altamira Cave, Spain

7.14. The chronology of major late Paleolithic technological innovations in Eurasia

7.15. Approximate locations of Siberian Mousterian and Upper Paleolithic sites

7.16. Structural ruins at the Mezin Upper Paleolithic site, Ukraine

7.17. Structural ruins at the Mezhirich Upper Paleolithic site, Ukraine

7.18. Structural ruins at the Pushkari 1 Upper Paleolithic site, Ukraine

7.19. Grooved antler artifacts from the Korolevo I Siberian Upper Paleolithic site

7.20. Use of the spear-thrower and a decorated example from southern France

7.21. Bone artifacts from the Later Stone Age layers of Nelson Bay Cave, South Africa

7.22. Approximate locations of key decorated caves in France and Spain and of caves in Germany with some of the oldest known ivory figurines

7.23. Mammoth ivory lion-man from an early Aurignacian layer at Holhenstein-Stadel, Germany

7.24. The early Aurignacian Venus from Galgenberg Hill, near Krems, Austria

7.25. Approximate locations of the Upper Paleolithic Gravettian sites with female figurines or engravings

7.26. Mammoth ivory Venus figurines from Kostenki 1, layer 1, Russia

7.27. Plan of a burial pit dated to roughly 30 ka at Kostenki XV on the Don River, Russia

7.28. Approximate locations of major Paleolithic sites in eastern Europe

7.29. Human figurines from the Mal’ta Upper Paleolithic site, Siberia

7.30. Structural remnants at the Mal’ta Upper Paleolithic site, Siberia

7.31. Grave with a fragmentary child’s skeleton at the Mal’ta Upper Paleolithic site, Siberia

7.32. Northeastern Siberia and northwestern North America at the height of the Last Glaciation

7.33. The main in situ occurrences of Clovis Paleo-Indian artifacts and of like-aged archaeological sites in the Americas

7.34. Map of Sunda and Sahul at the height of the Last Glaciation

8.1. A working phylogeny of the hominins from 4.5 million years ago to the present

8.2. Artifact types that characterize major Stone Age culture-stratigraphic units

8.3. Hominin Encephalization and Megadontia Quotients versus time

TABLES

1.1. A classification of living people at twenty-two potential levels in the Linnaean hierarchy

2.1. Basic parameters of some isotopes used in numerical dating

3.1. A provisional classification of the Primates to the level of the suborder and to the level of the family within the suborder Anthropoidea

3.2. Subdivision of the Hominoidea, with special reference to the Hominidae

4.1. Sedimentary units with hominoid fossils, artifacts, or both in the Tugen Hills

4.2. Hominin fossils from Lothagam

4.3. Hominin taxa and artifact traditions in the Koobi Fora Formation

4.4. Hominin taxa in the Lower Omo River Basin

4.5. Hominin taxa and artifact traditions in the Nachukui Formation

4.6. Hominin fossil localities in the Middle Awash region

4.7. The earliest putative hominins

4.8. African Pliocene sites with large fossil samples and no hominins

4.9. The principal craniums and mandibles from which the evolutionary relationships of the australopiths and early Homo must be inferred

4.10. Body, brain, and dental size estimates for extant Pan troglodytes, Homo sapiens, and various fossil hominins

4.11. Sites with fossils of Homo that antedate 1.9–1.8 million years ago

4.12. The main sites assigned to the Oldowan Industrial Complex

5.1. The main Javan Homo erectus sites

5.2. The main Chinese Homo erectus sites

5.3. African sites with fossils of Homo ergaster

5.4. African sites with fossils of early Homo sapiens

5.5. European sites with fossils of early Homo neanderthalensis

5.6. Endocranial capacity in Homo ergaster and later fossil representatives of Homo

5.7. The minimum numbers of postcranial elements recovered from the Sima de los Huesos, Atapuerca, through 1995

5.8. Characters that fossils from the Sima de los Huesos, Atapuerca, share with fossils of Homo erectus, the classic Neanderthals, and H. sapiens

5.9. Sites with fossils that could document late Homo erectus or H. heidelbergensis in China

5.10. European sites that have provided only doubtful evidence for human presence before 600,000 years ago

5.11. The main sites that document human presence in Europe before Global Isotope Stage 5

5.12. Early west Asian archaeological and human fossil sites

5.13. Key African archaeological sites where Homo ergaster, H. heidelbergensis, or early H. sapiens probably made the artifacts

5.14. The principal east and central Asian early archaeological sites

6.1. European sites that have produced important Neanderthal fossils since 1920

6.2. The African contemporaries of the Neanderthals

6.3. African sites where numeric dating indicates the Middle Stone Age (MSA) began before 250–200 ka

6.4. The principal west and central Asian Mousterian sites that have provided Neanderthal or near-modern human remains

6.5. The principal stratified MSA sites in southern Africa

6.6. East African sites that were occupied in the gap between 60 and 30 ka when southern and northern Africa seem to have been largely abandoned

6.7. Mousterian/MSA sites with wooden artifacts

6.8. Mousterian/MSA sites with proposed art or personal ornaments

6.9. Prominent open-air Mousterian and MSA sites in Europe, western Asia, and Africa

6.10. Mousterian/MSA sites that may preserve remains of structures

6.11. Mousterian/MSA sites that have provided remains of possible or probable food plants

6.12. South African sites that illuminate Middle and Later Stone Age coastal foraging

6.13. North African, European, and west Asian coastal or near-coastal sites to which Mousterian/MSA people brought shells of edible species

7.1. Non-European sites, in order of discovery, that have provided especially informative fossils of early fully modern humans

7.2 The principal African sites that have provided modern or near-modern human fossils older than 50 ka

7.3. Sites with Neanderthal bones that have provided mitochondrial DNA

7.4. African sites with ostrich eggshell beads that antedate 30 ka

7.5. Late Paleolithic sites with remains of plants that people probably introduced

PREFACE TO THE THIRD EDITION

Paleoanthropology continues to prosper, and the broad pattern of human evolution is becoming ever clearer, even if many subsidiary issues remain to be settled. This book summarizes the overall pattern as a series of five phases: (1) an australopithecine (or australopith) phase from 4.5 Ma (million years ago) or before until roughly 2.5 Ma; (2) a phase initiated by the appearance of the genus Homo and the first archaeological sites about 2.5 Ma; (3) a phase marked by the emergence of Homo ergaster (or early African Homo erectus) 1.8–1.7 Ma and the first dispersal of humans from Africa; (4) a phase initiated by an abrupt increase in brain size to near the modern average roughly 700–600 ka (thousands of years ago), the concomitant appearance of more sophisticated stone artifacts in Africa, and a subsequent Out-of-Africa event that produced the first permanent settlement of Europe; and (5) a phase beginning about 50 ka when modern humans spread from Africa to swamp or replace nonmodern Eurasians. The phase scheme is a heuristic device rooted in chronological change, but the fossil record itself is patently more complex. After 3 Ma, it involved repeated branching events that brought forth contemporaneous hominin (human) species. By 50 ka there were at least three nonoverlapping species, Homo sapiens in Africa, Homo neanderthalensis in Eurasia, and Homo erectus in the Far East. It was of course Homo sapiens that spread from Africa 50 ka to replace the others.

This book retains the fundamental structure of its predecessor, but each chapter has been substantially revised and updated. The first chapter outlines current understanding of the evolutionary process. The second addresses the geologic time frame for human evolution, with emphasis on numeric dating methods and on climatic change as a major driving (natural selective) factor. It devotes roughly equal space to the radiocarbon, radiopotassium, and luminescence dating methods, although the reliability of the luminescence methods is often questionable. Their main strength is their applicability to materials and time ranges that neither radiocarbon nor radiopotassium cover.

The third chapter summarizes primate evolution before the emergence of hominins roughly 7 Ma. It is the only chapter whose length has been reduced from prior editions. The reason is that as the prehominin primate fossil record has grown, so have professional disagreements on how to interpret it, and it has become increasingly difficult to synthesize succinctly and to relate directly to human evolution.

Chapters 4–7 present the five basic phases of human evolution listed above, together with the branching phylogenetic tree that crosscuts them. These chapters are longer than their counterparts in the previous edition, mainly because they incorporate the implications of important new discoveries. The new findings include fossils dated between 7 and 5 Ma that may document the oldest hominin species; fossils dated to 2.5 Ma that shed fresh light on the australopithecine roots of Homo; sites that may imply people reached northern China as much as 1.6 Ma; other sites that suggest people sporadically penetrated southern Europe between 1.1 Ma and 800 ka and northern Europe by perhaps 700 ka; new dates and additional fossils to document the evolution of Homo neanderthalensis in Europe between 500 and 70 ka; additional dates and fossils that illuminate the evolution of Homo sapiens in Africa over the same interval; and finally, the ongoing retrieval of Neanderthal DNA, which ever more strongly underscores the divergence of the Neanderthals from modern humans. Like all overviews of human evolution, the one presented here is a narrative, but it is constrained by evidence, and the inclusion of fresh evidence and analyses means that it approximates the actual course of human evolution more closely than its predecessors.

Chapter 8 summarizes the main points in chapters 4–7. It is a synopsis or abstract that could be read first as well as last.

Not all the most recent findings are equally persuasive, and in some cases, including for example, the settlement of northern China by 1.6 Ma and of northern Europe by 700 ka, they require confirmation from additional discoveries. This follows from the realization that the fossil and archaeological records are inherently noisy and that multiple well-substantiated observations are required to identify a valid signal. My own belief, repeatedly expressed in the text, is that a first occurrence should be treated as a possible accident and even a second should be regarded as a possible coincidence. Only repeated, independent, mutually consistent occurrences can document a reliable pattern. This philosophy explains what some may find surprising—I have addressed the hobbit (Homo floresiensis) only in passing, even though it may one day bear out its frequent billing as one of the two or three most important fossil finds of the past hundred years.

The recurring shortage of reliable, mutually supportive observations largely explains what lay observers and science writers often see as the contentious nature of paleoanthropology, in which practitioners stake out contrary positions that seem irresolvable. Some issues may be truly irresolvable, but many others require only fresh observations to reduce or eliminate ambiguity, and the problem is that new observations often appear slowly. To see real progress, we must often then be willing to wait years or even decades. The Neanderthals provide a case in point. As recently as twenty years ago, some prominent specialists argued that the Neanderthals were directly ancestral to modern Europeans or that we could never know their evolutionary relationships, for their bones told us only about their behavior. Today, thanks to fresh fossil finds and dates, to multiple, consistent analyses of the genes of living humans, and perhaps above all, to the spectacular recovery of DNA from Neanderthal bones, almost all specialists agree that the Neanderthals represent an evolutionary dead end. Many authorities who formerly placed the Neanderthals on the line to modern Europeans now argue only that they interbred with modern human invaders. However, genes increasingly deny even this fallback position, and the bottom line is that the Neanderthal controversy, which had raged for over a century, is now essentially concluded. This allows us to concentrate on other questions like the reason for their demise. Specialists disagree sharply on this issue, but as laid out in this book, genes again have the power to narrow the alternatives.

Like the previous editions, this book differs from most commercial texts on human evolution in the extensive integration of the fossil and archaeological records and in the amount of supporting detail for both. The quantity of information in the previous editions makes me suspect that they were used more as references than as texts. In an attempt to enhance the value of this version as a readable text, I have removed much of the detailed information to tables, which I believe makes for smoother reading and easier access. I’ve also removed all bibliographic citations to separate paragraphs that immediately follow each topic within each chapter so that students can treat the citations as optional. An alternative scheme, which I used in the second edition, was to replace conventional in-text author-date citations with numbered references, but professionals disliked having to turn to the bibliography to see who was being cited. I hope both students and professionals will welcome the present system.

The literature on human evolution is mushrooming, and this edition contains about 1,700 more references than its predecessor. The new references appeared mostly in the last ten years, and they represent only those whose titles or abstracts suggested that I needed to read them. They comprise only a fraction of the articles and books I might have read, and I apologize to anyone who produced an important source that I inadvertently overlooked.

Finally, I’m grateful to readers who pointed out errors or omissions in the previous editions, and I hope they find I have made appropriate corrections. I owe a special debt to Kathryn Cruz-Uribe, who illustrated most of the fossils and artifacts in the earlier editions and who produced fifty new high-quality illustrations for this one. I thank David DeGusta, Teresa Steele, and Tim Weaver for thoughtful comments on parts of the present text.

PREFACE TO THE SECOND EDITION

Paleoanthropology—the parent discipline for human paleontology and paleolithic archeology—has never been more vigorous or productive. In the ten years since the first edition of this book was published, paleoanthropologists have discovered human fossils that antedate 4 million years ago and that show ever more clearly the ape origins of the human family; they have found new fossils and archeological sites that imply a complex history of branching events within the genus Homo; and they have uncovered a wealth of new fossil, archeological, and genetic evidence that the last shared ancestor of all living humans existed in Africa no more than 200,000 years ago.

The purpose of this edition is to show how the new discoveries and analyses supplement previous ones to reveal the basic course of human evolution. Two conclusions are especially clear. The first is that the australopithecines and other very early people who lived between 5 and 1.8 million years ago were intermediate in appearance and behavior between apes and unquestionable humans and that the first true humans, who appeared 1.8 to 1.7 million years ago, were morphologically and behaviorally intermediate between the australopithecines and living people. An ironic corollary is that the human family now provides one of the most persuasive fossil cases for the occurrence of macroevolution.

The second major conclusion is that fresh fossil, archeological, genetic, and geochronological findings confirm earlier ones, suggesting that modern humans originated in Africa and later replaced other kinds of people in Eurasia. The first edition espoused this Out-of-Africa scenario only cautiously, and it retained an earlier view that human evolution comprised a series of grades or stages, from the relatively apelike australopithecines through Homo habilis, Homo erectus, and early (or archaic) Homo sapiens to modern Homo sapiens.

The richer fossil, archeological, and genetic records that bolster Out of Africa now indicate that the stage system must be discarded in favor of a branching scheme. Thus the text argues that after an initial human dispersal from Africa by 1 million years ago, at least three geographically distinct human lineages emerged. These culminated in three separate species: Homo sapiens in Africa, Homo neanderthalensis in Europe, and Homo erectus in eastern Asia. Homo sapiens then spread from Africa beginning perhaps 50,000 years ago to extinguish or swamp its archaic Eurasian contemporaries. The spread was prompted by the development of the uniquely modern ability to innovate and to manipulate culture in adaptation. This ability may have followed on a neural transformation or on social and technological changes among Africans who already had modern brains. Whichever alternative is favored, the fossil, archeological, and genetic data now show that African H. sapiens largely or wholly replaced European H. neanderthalensis. The situation in eastern Asia is more obscure, because the fossil, archeological, and geochronological data are much sparser, but fresh data will probably show that H. sapiens replaced H. erectus in the same way.

The change from a synthesis centered on stages to one based on branches required a new chapter titled "Evolution of the Genus Homo in place of the first edition’s separate chapters titled Homo erectus and Early Homo sapiens." The new chapter stresses the fossil and archeological evidence for multiple, contemporaneous species within fossil Homo, but it also acknowledges the continuing meagerness of the record, particularly before the advent of unequivocal Homo sapiens and Homo neanderthalensis by 130,000 years ago. To ensure that readers appreciate just what the evidence is (and what it is not), the new chapter illustrates many of the key fossils and artifacts.

Most of the remaining chapters retain their original titles, but every one has been updated and expanded, and they all have many new illustrations. The total number of figures for the book has been increased from 151 to 223, and I could not have produced them without the willing, skilled assistance of my archeological colleague and friend Kathryn Cruz-Uribe. Even a cursory examination of the following pages will reveal her enviable ability to render informative, tasteful line drawings of complex fossils and artifacts. I used the computer programs Adobe Streamline and Macromedia Freehand to enhance her drawings with labels for key landmarks or features.

Like the first edition, this one differs from most commercial texts in emphasizing not only the broad pattern of human evolution but also the fossil and archeological data behind it. The book includes far more fossil and archeological detail than standard introductions to human evolution, and it is intended to be as much a sourcebook as a text. Nonetheless, the first edition was often used as a text, and with this in mind I have tried to make the second edition more accessible to undergraduate students. The most significant change is the addition of an introductory chapter that outlines how species evolve and how evolutionary biologists (including paleoanthropologists) classify and name them.

My own students have often complained about the standard academic author-date referencing system used in the first edition, because it obstructs text flow and intelligibility, especially when many citations are chained together. In this edition, I have therefore adopted a system in which numbers in parentheses in the text refer to numbered items in a summary reference list. A secondary benefit of this system is that it shortens the text by more than five thousand words. The reference list is alphabetized, so works by a particular author are easy to locate. There is also a separate reference index that lists the text pages on which individual numbered works are cited. I used the Endnote computer program to ensure that the numbers cited in the text matched the numbered works in the reference list.

The reference list includes more than 2,500 separate items, yet this is only a tiny fraction of the literature on human evolution. It is in fact only a small fraction of the items that have appeared since the first edition was published, and it may neglect some significant discovery or idea in a reference I have missed. I hope that anyone who sees a glaring omission will notify me so I will not overlook it in the future.

In preparing the second edition, I have benefited from the research and writing of innumerable colleagues, but I particularly want to thank James Bischoff, Frank Brown, Kathryn Cruz-Uribe, Robert Franciscus, Clark Howell, and Henry McHenry for critical comments on portions of the text. I am also grateful to the many wonderful students at the University of Chicago and Stanford University who endured the lectures and text drafts that underlie both editions.

PREFACE TO THE FIRST EDITION

Superficially, the study of human evolution seems remarkably abstruse and impractical. Yet, each year the popular press covers major finds, large prime-time audiences watch televised documentaries, and thousands of students enroll in pertinent university courses. Australopithecus and Olduvai Gorge feature in nationally syndicated cartoons, and some discoverers of important fossils are better known than their counterparts in more practical fields such as physics and medicine. Clearly, in spite of its apparent irrelevance to everyday affairs, the origin of the human species is intensely interesting to most modern humans, who are fascinated by the growing number of fossils, artifacts, and related facts that scientists have amassed. They want to know what these data tell us about the appearance and behavior of our remote ancestors. This book is a summary of what I think the data say.

There are many perspectives on how the data should be interpreted, and this book, of necessity, reflects just one. In writing it, I have tried to steer a middle course between what I see as two extreme approaches—one in which the data are simply a springboard for stimulating speculation about what might have happened in the past and another in which they are meaningless except to test and eliminate all but one competing explanation of what really happened. The difficulty with the first perspective is that it emphasizes imagination over validity. The difficulty with the second, whose roots lie in a perception of how the physical sciences have advanced, is that it assumes an unrealistic degree of control over data quantity and quality. In fact, substantial control is unusual in human evolutionary studies, where carefully planned experiments are rare and most data are obtained through excavations and field surveys whose success often depends more on chance than on design. Under these circumstances, I think that the physical sciences provide a less suitable role model than the judicial system, in which often limited evidence is weighed to determine which of two or more competing explanations or interpretations seems most reasonable. In most instances the possible alternatives are not pared unequivocally to one, but one is selected because it seems more justifiable, given the evidence on hand.

Of course, in human evolutionary studies, as in the judicial system, both laypeople and specialists may disagree about what is reasonable or justifiable and also about the soundness of the supporting evidence. All too often the evidence is incomplete, ambiguous, or even contradictory, and it cannot be used to bolster any particular theory or explanation very strongly. In this book I have tried hard to present the major competing opinions on prominent unresolved issues, and whenever possible I have explained why I think one view is more reasonable than others. More often than I would like, I have had to say that a firmer choice will require more data. I know that not everyone will agree with the positions I have taken or even with my decision to abstain on some matters. However, I think that such differences of opinion are unavoidable, given the imperfect nature of the evidence; and the point is that this book is inevitably just one of many possible summaries of what we know about human evolution. Its success will depend on the extent to which the readers, experts, and laypeople alike think that the presentation and argumentation are sensible.

Philosophical approaches aside, there are several possible ways to organize the evidence for human evolution. The way I have chosen is perhaps the most conventional, focusing on a series of chronologically successive stages—beginning with the earliest Primates, dating from perhaps 80 million years ago, and ending with the emergence of anatomically modern people within the past 200,000 years. The presentation does depart from the norm, however, in that I have given roughly equal weight to the fossil record and to the accompanying archeological evidence over the 2.5 million year interval for which this is available. Most summaries focus largely on the fossils or on the archeology, thereby forgoing one of the major points that I have sought to make—namely, that the human form and human behavior have evolved together and that neither can be fully understood or appreciated without a full understanding of the other. At the same time, however, it remains true that the fossils are far easier to arrange into a set of chronologically successive, interrelatable units, and, since the fossil record is also far longer than the archeological one, I have relied on the fossils to define the chronologically successive stages that structure the text. This is not to say that there are no problems in defining the fossil units, but these pale beside the difficulties in defining and interrelating corresponding archeological categories. The difference stems from our much weaker understanding of the mechanisms underlying artifactual (cultural) change and differentiation.

A second way in which this survey differs from many others is that it includes more information on specific sites, fossils, artifacts, and so forth—in short, it is more detailed, with more concern for the factual evidence that underlies our understanding of human origins. It is a formal rendering of the lectures I give in an upper-level undergraduate course at the University of Chicago, and it has been written with upper-level undergraduates, graduate students, and professionals in mind. In my experience, the audience for whom it is primarily intended will already have a basic understanding of how evolution occurs (through natural selection, mutation, gene flow, and gene drift), and therefore this latter topic is not explicitly addressed. However, at least some members of the audience will lack essential background information on skeletal anatomy, zoological classification and nomenclature, stone tool typology and technology, and especially the geologic time frame for human evolution, and so these subjects are covered. In general I think the book is too detailed to be a central text in lower-level courses, especially ones that also deal with modern human variation, genetics, and the like; but I hope it will find use there as one of the sources the instructor consults or recommends to those students who are especially curious about the fossil and archeological records.

In keeping with the comparatively technical orientation of the book, I have employed an in-text citation system that is common in professional scientific publications. I rejected the usual textbook system of grouping sources at the end of each major section or chapter because I felt the target audience for the book would prefer to know precisely where to look for further information on a particular topic. I also wanted to give credit directly where it was due. I rejected a system of linking references to numbers because I thought the risk of serious error would be too great when so many references are involved. The large number of references was unavoidable, given the broad theme, but I have tried to keep the list manageable by stressing recent sources that can serve as guides to older ones, and I have also excluded many non-English primary sources in favor of secondary English ones with their own extensive bibliographies of important non-English publications.

No synthesis of human evolution would be successful without good illustrations, but these can be very expensive and time consuming to produce. As a result, even many commercially produced texts are underillustrated. I have attempted to compensate for the limitations of time and expense that were important here by adapting many illustrations from published sources, which are gratefully acknowledged. Thanks mainly to the efforts of Kathryn Cruz-Uribe, most of the illustrations have been substantially modified to support pertinent points in the text and to provide stylistic consistency. In addition, whenever possible, I have labeled important features directly on fossils, artifacts, site plans, and so forth, and I have attempted to make the captions freestanding supplements to the text, to emulate the useful sidebars that are common in commercially produced texts. My goal was to make the illustrations especially helpful to those with little or no prior knowledge of skeletal anatomy, stone artifacts, stratigraphies, and such.

It was not easy to choose a title for the book, because the most obvious ones, such as Human Origins and Human Evolution, have been used many times before. The final choice—The Human Career—is the name of a graduate course I took at the University of Chicago in 1962, in which F. Clark Howell introduced me to the concept of human evolutionary studies as an amalgam of human paleontology and paleolithic archeology. Howell’s alternative name for the subject matter, paleoanthropology, would do equally well—though it too has been used before and has often been applied to human paleontology alone rather than to the broader paleontological/archeological field that Howell had in mind. In both the title and the text, I intend the vernacular term human (and its complement, people) to refer to all members of the zoological family Hominidae, as conventionally defined, and not simply to living humans.

My own research on human evolution has focused mostly on behavioral (archeological) evidence from middle and late Quaternary sites in southern Africa and parts of Europe, and my acquaintance with the remainder of the record comes mainly from published sources. In synthesizing these, I have tried hard to make the text as accurate, comprehensive, and up-to-date as possible, and I have been greatly helped by comments and criticisms from Peter Andrews, Kathryn Cruz-Uribe, Janette Deacon, Leslie Freeman, Fred Grine, Clark Howell, Philip Rightmire, Chris Stringer, Russell Tuttle, and Tom Volman. I hope they find that I have employed their suggestions productively and that I have not introduced any new errors in the process. Inevitably, however, some defects remain, and I would be grateful to hear from anyone who finds a specific problem or who has suggestions on how the interpretations or overall organization can be improved.

1

Evolution, Classification, and Nomenclature

This chapter introduces basic evolutionary terms and concepts that are essential if we are to synthesize sites, fossils, artifacts, dates, and other facts into a coherent overview of human evolution. The discussion draws on much more extensive treatments of process and principle in works listed immediately below.

GENERAL SOURCES: Cain 1960; Dawkins 1987; Dobzhansky 1962; Eldredge 1991, 1995; Gould 2002; Levinton 1988; Maynard Smith 1989; Mayr 1942, 1963, 2001; Ridley 1986, 1993; Simpson 1953, 1961; Stanley 1979, 1981

The Biological Species

The species is the least arbitrary and most fundamental evolutionary unit, and it must be understood before any consideration of evolution, even one focused tightly on a single species like Homo sapiens. Evolutionary biologists define a species as a group (or population) of organisms that look more or less alike and that interbreed to produce fertile offspring. This definition is often called the biological species concept. In practice, individuals are usually assigned to a species based on their appearance, but it is their membership in the same procreative unit that validates (or invalidates) the assignment. Thus, no matter how detailed the resemblances between two groups of organisms, if individuals cannot exchange genes between groups, the two populations must be assigned to different species.

Species ordinarily comprise lesser units known as breeding populations (sometimes called demes or Mendelian populations) in which most individuals find their mates. For example, the American black bear species (Ursus americanus) ranges across much of North America, but black bears in Alaska and central Mexico are unlikely to mate, and they are thus reasonably assigned to separate breeding populations. The maximum breeding population is always the species, since no members may find mates outside it. Mountain chains, deserts, large water bodies, or other barriers between breeding populations may prevent them from interbreeding, but as long as interbreeding is possible and fertile offspring would result, the breeding populations still belong to the same species. Breeding populations tend to fluctuate in size, membership, and total number over relatively short periods, which means they are harder to delineate than species. However, as discussed below, they are central to evolution, since they are potential species in the making.

The modern definition of the species, founded in interbreeding capacity, is not the only conceivable one, and it is worthwhile to consider why it has replaced a definition that was popular before evolution became widely accepted in the second half of the nineteenth century. This earlier concept defined each species according to the characters of a type specimen, usually the first one to be found or described. Specimens that were not identical to the type specimen were allowed in the same species, but they were regarded as sports or deviants, and their anomalous features did not contribute to the species description.

The typological definition of the species was problematic even at the time it was most popular. From the beginning, its arbitrariness was apparent, and it floundered when additional specimens turned out to typify the species better than the original type. It was also difficult to decide just how different two specimens had to be before they could be assigned to different species. Typological practice tended to exaggerate small differences, and the result was an excessive number of species. Most important, however, the typological definition literally fixed species in time. Species were immutable, for if they could change then there could be no type specimen, only an endless series of deviants. The typological definition was therefore incompatible with organic evolution, and the infraspecific (within-species) variation it explicitly suppressed was fundamental to Darwin’s novel idea of how species evolved, via natural selection of the fittest (discussed below).

Only the modern biological species definition is consistent with the idea of evolution, but ironically, only the typological definition is directly applicable to the fossil record. The disjunct arises partly because interbreeding cannot be observed among fossils, and paleontologists must rely strictly on morphological features. In practice, with the biological species concept in mind, they commonly use skeletal variability within living species to evaluate whether differences among fossils are likely to mean interbreeding could not occur. In this case the fossils are assigned to different species. However, the degree of skeletal variability varies from species to species, which means there can be no universal standard. Sometimes, as in the case of the African guenon monkeys (Cercopithecus), even members of different species are difficult or impossible to distinguish from skeletons alone. In other cases, for example, modern Homo sapiens, substantial skeletal variation occurs within a single unquestioned biological species. Paleontologists face the additional difficulty that their specimens are usually not complete skeletons. Isolated bones or fragments of bones are much more common, and this helps to explain why fossil species are often so hotly debated.

Finally, there is the problem that the identification of fossil species inevitably requires dividing the evolutionary continuum into arbitrary units. If the record is complete enough, some dividing lines will fall between succeeding generations that could have interbred and that therefore should not be assigned to different species. This problem seems insuperable, except for the observation that whereas evolution is continuous, the fossil record often is not. As discussed below, the fossil history of most species suggests long periods of morphological stability or stasis, followed either by extinction or by brief bursts of rapid change during which new species emerge. In this circumstance, the chances of finding fossils from populations in the process of speciation are slim, and the frequent absence of such fossils therefore poses no theoretical problem for the biological species concept. Their absence means that most fossil species, therefore, have discrete beginning and endpoints that probably correspond at least broadly to the reproductive boundaries of living species.

Natural Selection

Nonspecialists often think that Charles Darwin (1809–1882) devised the concept of evolution in The Origin of Species (published in 1859). In fact, others had proposed the idea long before, and Darwin’s monumental contribution was the discovery of a natural mechanism to explain how evolution had occurred. Before Darwin, the notion that species had evolved was hardly less mystical than the more common traditional idea that they had been created supernaturally.

Darwin’s great discovery was the principle of natural selection, which he synthesized from his own natural history observations and from the writings of other eminent nineteenth-century thinkers, perhaps above all Thomas Malthus (1766–1834) on population growth. As framed by Darwin and still employed by evolutionary biologists, natural selection is grounded in three observations: (1) that individuals within a breeding population vary in morphology and behavior, (2) that offspring tend to inherit features of morphology and behavior from their parents, and (3) that not all individuals contribute equal numbers of offspring to the next generation. If we assume that differential capacity to survive and reproduce is linked to morphology and behavior, then any novel traits that enhance survival and reproduction will tend to increase in frequency from one generation to the next. Such traits are said to be adaptive or in the parlance of evolutionary biology, to make their bearers more fit, and natural selection can be defined as the sum of environmental forces or pressures that determine fitness. Fitness is defined totally, if somewhat tautologically, as an individual’s ability to produce offspring who themselves survive and reproduce.

As Darwin noted, natural selection is exemplified by the well-known process that people have used to produce breeds of domestic animals. In this case the selective force is human preference or desire, and an individual animal’s fitness (ability to survive and reproduce) depends on the extent to which it exhibits desired characteristics. New breeds can be produced relatively quickly, even within historical memory, if selection is stringent enough. Darwin recognized that far more time would be required to transform one species into another or to produce two species from one, and he probably would not have proposed natural selection as the underlying mechanism if geologists had not simultaneously demonstrated the great antiquity of the earth.

Darwin followed contemporary geological research closely, and he was especially influenced by the work of Charles Lyell (1797–1875), often known as the founder of modern geology and the formulator of the uniformitarian principle that guides it. In its simplest form, uniformitarianism is the notion that the present is the key to the past. Less cryptically, it is the idea that erosion, sedimentation, volcanism, crustal movements, and other geologic processes that can be observed today have operated throughout earth history and are sufficient to explain the geologic record. From a uniformitarian perspective, for example, crustal uplift can explain the occurrence of shelly marine limestone far above modern sea level. To accept this, an observer must accept only that uplift and other geologic processes that produce only small-scale effects over the short term can produce much greater large-scale effects over eons. In essence, Darwin’s idea that natural selection explains the origin of species simply extended the uniformitarian principle to the biological realm.

Genetics and Evolution

Darwin believed that natural selection operated on individuals, and fitness in the classical Darwinian sense applies exclusively to individuals. But modern evolutionary biologists often think of selection as operating on particular traits. The difference is profound, because it reflects an understanding of how traits are inherited—in short, of genetics—that was totally lacking in Darwin’s day. Strictly speaking, the fundamental nature of inheritance had been established, but the discoverer, the Austrian priest Gregor Mendel (1822–1884), published his findings in an obscure journal and his pioneering research was recognized only in 1900, long after both he and Darwin had died.

When Darwin wrote The Origin of Species, it was generally believed that offspring blended the traits they inherited from their parents and they then passed on a blend of blends to their own offspring. This view of inheritance presented an immediate problem for the idea of natural selection, for it followed that a selectively advantageous novelty (or mutation) could not become fixed in a population. Instead, it would become diluted or lost through progressive blending with the much larger number of less advantageous alternatives. In essence, Mendel showed that inheritance was not a blending process but, instead, involved discrete units or particles, inherited half from one parent and half from the other. The particles, which we now call genes, retain their integrity from generation to generation, even though their expression may vary depending on the presence of other genes or on particular environmental conditions. Thus an adaptive or advantageous novelty will not necessarily be diluted out of existence but could spread if it confers greater fitness.

Today a particular gene may be defined by its position on a chromosome (its locus) or by its singular chemical structure. Evolutionary divergence

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